QL 

368 

F6C83 


SMITHSONIAN  INSTITUTION 

UNITED  STATES  NATIONAL  MUSEUM 
Bulletin  100 

VOLUME  1,  PART  6 


CONTRIBUTIONS  TO  THE  BIOLOGY  OF  THE 

PHILIPPINE  ARCHIPELAGO  AND 

ADJACENT  REGIONS 


THE  RELATIONSHIPS  OF  THE  GENERA 

CALCARINA,  TINOPORUS,  AND  BACU- 

LOGYPSINA  AS  INDICATED   BY 

RECENT  PHILIPPINE  MATERIAL 


JOSEPH  A.  CUSHMAN 
Of  the  Boston  Society  of  Natural  History 


WASHINGTON 

(;OVI:RNMI:NT  PRINTING  OFFICK 
1919 


LIBRARY 
UNIVERSITY  OF 

SANTA  BARBARA 


PL 


THE  RELATIONSHIPS  OF  THE  GENERA  CALCARINA, 
TINOPORUS,  AND  BACULOGYPSINA  AS  INDICATED 
BY  RECENT  PHILIPPINE  MATERIAL. 


By  JOSEPH  A.  CTTSHMAN, 

Of  the  Boston  Society  of  Natural  History. 


During  the  Albatross  Philippine  Expedition  of  the  United  States 
Bureau  of  Fisheries  great  numbers  of  shallow- water  foraminifera 
were  collected.  Those  belonging  to  Calcarina  and  Tinoporus,  as 
those  genera  are  usually  understood,  form  a  considerable  amount  of 
material  from  many  stations  and  hundreds  of  specimens.  The  prob- 
lem of  identifying  the  species  represented  has  not  been  a  simple 
problem  and  has  involved  a  review  of  much  of  the  earlier  literature. 
Much  of  the  difficulty  of  the  problem  has  centered  about  the  question 
which  other  workers  have  had  as  to  the  exact  identity  of  Montfort's 
genus  Tinoporus. 

An  indication  first  of  the  various  species  involved  and  later  their 
generic  position  will  perhaps  be  the  easiest  way  to  present  the  results. 
The  earliest  species  is  the  Nautilus  spengleri  Gmelin,  not  Linnaeus,  as 
usually  given,  as  this  first  appears  in  the  thirteenth  edition,  1788, 
which  is  Gmelin's,  based  on  the  figure  given  by  Spengler  in  1781  as 
"  Ammonshorn."  This  species  is  now  apparently  well  defined.  D'Or- 
bigny  in  1826  referred  to  it  under  his  genus  Calcarina  spengleri 
Gmelin  and  gives  as  synonymous  Tinoporus  baculatus  Montfort  and 
Siderolites  calcitrapoides  Lamarck.  No  figures  are  given,  but  those 
in  the  "  Planches  inedites,"  published  by  Fornasini,  include  all  of 
d'Orbigny's  species  of  the  1826  paper.  The  figures  given  in  the 
Challenger  Report,  H.  B.  Brady,  1884,  give  a  good  idea  of  the  species. 
It  is  a  lenticular  test,  biconvex,  made  up  of  numerous  chambers  in  a 
close  coiled  trochoid  spire,  developing  a  secondary  skeleton  and  with 
a  series  of  blunt  spinose  processes  about  the  margin  of  the  test  taking 
their  origin  early  in  the  development  of  the  test  and  gradually  in- 
creasing in  size.  The  surface  is  generally  smooth  or  somewhat  tuber- 
culate,  especially  in  the  center  of  the  disk  at  either  side,  and  the 
spiral  condition  continues  throughout  the  life  history.  The  spines 
are  smooth  except  for  the  channels  of  the  supplementary  canals,  and 
are  bluntly  rounded  at  the  extremities,  usually  from  three  to  six  or 
more  with  five  or  six  the  usual  number. 

121299-19  363 


364  BULLETIN  100,   UNITED  STATES  NATIONAL  MUSEUM. 

Such  tests  (pi.  44,  fig.  1)  are  fairly  common  in  the  Philippine  ma- 
terial, at  some  of  the  stations  being  very  common,  and  there  seems 
little  question  as  to  what  spengleri  Gmelin  really  is. 

Of  the  synonyms  given  by  d'Orbigny,  Siderolites  caldtrapoides 
Lamarck  is  based  upon  a  chalk  fossil  from  Maestricht  and  I  have  been 
able  to  study  material  from  Maestricht  kindly  sent  me  from  the 
United  States  National  Museum,  and  this  is  not  the  same,  as  will  be 
shown  later. 

The  next  problem  is  Tinoporus  baculatus  Montfort.  This  has  been 
the  cause  of  much  dissension.  Montfort  refers  to  an  earlier  figure 
of  Fichtel  and  Moll  of  1803  (pi.  15,  figs.  i.  &.),  Montfort  also  figures 
the  species.  This  figure  shows  a  rounded  test  with  three  truncate 
spinose  projections  at  three  of  four  equidistant  points  of  the  circum- 
ference, the  exterior  of  the  test  black  with  white  points  on  both  the 
body  and  spines.  The  interior  is  shown  as  spiral  [?]  and  composed 
of  numerous  chambers  with  an  indication  that  they  are  several  deep 
in  the  convex  area. 

In  the  Philippine  material  especially  from  Albatross  Station 
D5134  (Sulu  Archipelago,  near  Basilian  Island — Latitude  6°  44' 
45"  N.;  longitude  121°  48'  E—  25  fathoms),  there  are  numerous 
specimens  of  a  black  form  with  four  equidistant  spines  and  numer- 
ous raised  tubercles  over  the  surface,  all  corresponding  very  well 
with  the  crude  figure  given  by  Montfort.  The  short  generic  descrip- 
tion is  as  follows: 

"Coquille  libre,  univalve,  cloisonnee  et  cellulee,  spiree  et  lenti- 
culaire,  tet  granule  exterieurement ;  bouche  semi-lunaire,  placee  vers 
la  circonf  erence  et  sur  un  des  cotes  \  dos  carene,  arme  de  quatre  points 
au  plus ;  les  deux  centres  bombes  et  releves." 

The  specimens  from  D5134  and  elsewhere  (pi.  44,  fig.  3)  are  very 
clearly  the  same  as  those  Montfort  had.  It  is  a  test  very  similar  in 
shape  to  spengleri  Gmelin  but  has  a  greater  amount  of  secondary 
skeleton,  the  spines  covered  with  spinose  projections  (indicated  by 
the  white  dots  of  Montfort 's  figure)  and  the  whole  test  spinose  ex- 
cept the  centers  of  each  side,  which  have  high,  conical  tuberculations, 
and  on  the  ventral  side  the  last  formed  chambers  of  the  outer  whorl 
often  more  or  less  obscured  by  the  surface  ornamentation.  There 
are  usually  four  spines  at  equidistant  points  on  the  periphery,  some- 
times five.  This  seems  to  be  the  adult  of  this  particular  species, 
which,  as  has  been  said,  is  very  common  at  certain  stations.  Mont- 
fort's  specimens  were  from  the  East  Indies,  also  from  the  Arabian 
Sea  and  the  Adriatic.  The  sections  of  the  Philippine  specimens 
show  them  to  be  spiral  throughout,  with  a  certain  amount  of  piling 
up  of  chambers  in  the  central  portions  in  late  growth.  If,  then,  the 
baculatus  Montfort  is  taken  as  this  species,  a  second  definite  species 
is  segregated. 


CALCARINA,  TINOPOKUS,  AND  BACULOGYPSINA.  365 

D'Orbigny  in  1826  figures  Calcarina  defrancii  d'Orbigny  (pi. 
13,  figs.  5-7).  These  show  a  fairly  smooth  test  in  a  low  trochoid 
spire  with  elongate  spines,  one  from  each  chamber,  and  those  of  the 
earlier  whorl  persisting  above  those  of  the  last  formed  whorl.  Brady 
figures  similar  specimens  in  the  Challenger  report.  Fornasini  in 
figuring  the  tracings  of  the  "  planches  inedites  "  gives  more  bizarre 
forms,  in  one  with  spines  having  forked  tips.  Such  specimens  were 
figured  by  Carpenter  (Introd.  Foram.,  1862).  These  are  caused,  at 
least  in  some  cases,  by  the  spines  of  the  earlier  and  later  whorls  fus- 
ing, and  being  at  different  angles  the  points  of  the  two  or  more  con- 
tinue their  original  lines  and  diverge.  Specimens  of  this  species  are 
smooth  except  for  the  spire,  and  the  spines  are  also  relatively  smooth. 
Specimens  are  fairly  common  at  some  stations,  usually  in  compara- 
tively shallow  water  (pi.  44,  fig.  2). 

A  fourth  species  is  described  by  Brady  in  1876  and  figured  in  the 
Challenger  Report  (pi.  107,  figs.  8,  9)  as  Calcarina  hispida  H.  B. 
Brady.  This  again  is  a  well-known  species,  flattened  and  the  entire 
surface  hispid  with  short  blunt  spines,  these  extending  out  into  the 
numerous  flattened  spines  of  the  periphery.  This  spinose  condition 
is  part  of  the  supplementary  skelton  and  the  newly  added  chambers 
of  the  spire  are  added  directly  on  top  of  this  hispid  surface.  The 
chambers  themselves  at  their  inception  are  very  thin  walled  and 
punctate,  but  quickly  add  the  thickened  layer  of  spinose  skeleton. 
Occasionally  there  are  a  few  tubercles  developed  in  the  center  of  the 
surface  (pi.  44,  fig.  4) .  From  the  evidence  of  the  "  planches  inedites  " 
this  is  the  same  as  d'Orbigny's  Calcarina  quoyi,  which  becomes  a 
synonym  of  C.  hispida,  as  it  was  not  recognizable  until  the  publi- 
cation of  the  figures  by  Fornasini  in  1907. 

Associated  with  the  other  species  already  noted  were  specimens 
with  usually  three  rather  pointed  spines,  very  hispid,  the  center  of 
the  body  surface  with  a  group  of  large  tubercles  and  a  very  definitely 
trochoid  spire.  It  was  noted  that  these  occurred  only  in  association 
with  a  larger  form  of  what  has  been  generally  known  as  Tinoporus. 
A  series  of  these  studied  showed  that  all  were  but  developmental 
stages  of  one  species,  some  of  the  stages  of  which  are  shown  in  plate 
45,  figure  1,  shows  the  general  appearance  of  the  young,  with  a  regular 
spire,  the  elongate  spines  and  central  tubercles.  Figure  2  shows  a 
slightly  later  stage,  where  the  newly  added  chambers  now  begin  to 
appear  around  the  periphery  of  the  test  and  even  on  the  dorsal  side 
(26).  Figure  3  shows  a  later  stage  where  the  development  of  the 
chambers  has  become  greater  and  both  sides  are  beginning  to  be  cov- 
ered with  the  hemispherical,  thin  walled,  punctate  chambers  cover- 
ing the  hispid  surface  and  extending  out  onto  the  spines.  In  the 
adult  this  continues  until  the  spines  are  completely  covered,  as  in 
figure  5.  In  figure  4  is  shown  a  specimen  with  an  eroded  test  the 


366  BULLETIN   100,  UNITED  STATES-  NATIONAL  MUSEUM. 

spines  large  and  blunt,  and  at  x  the  remains  of  some  of  the  tubercles 
of  the  test  which  grow  outward  and  help  support  the  test  which  is 
otherwise  very  fragile.  This  gives  a  large  globular  test  in  the  adult 
(pi.  44,  fig.  5),  from  which  usually  project  three  spine  tips,  still  hispid 
if  not  covered  by  the  hemispherical  chambers,  the  chambers  large  and 
rather  coarsely  punctate.  This  species  seems  to  be  undescribed  and 
will  be  referred  to  later. 

Another  species  (pi.  44,  fig.  6),  not  common  in  the  Philippine  ma- 
terial but  very  abundant  in  the  Murray  Island  region  of  the  Great 
Barrier  Reef  of  Australia  and  elsewhere,  is  that  which  is  figured  by 
Brady  and  others  as  Tinoporus  baculatus.  This  species,  as  shown 
by  Carpenter  in  1860,  has  a  spiral  young,  bu£  the  spire  is  continued 
for  but  a  single  whorl  when  the  several  spines  are  produced,  and 
later  growth  is  on  the  order  of  Gypsina,  covering  the  test  with  con- 
centric layers  of  small  chambers,  interspersed  with  which  are  bosses 
of  clair  solid  shell  material  regularly  placed  and  connected  radially 
with  each  other.  The  spines  are  not  hispid  but  smooth  or  channeled 
and  are  usually  four  to  eight  or  nine  in  number.  The  chambers  are 
much  smaller  than  those  of  the  preceding  species,  and  not  so  ob- 
viously punctate,  while  the  reticulate  pattern  caused  by  the  bosses 
and  their  radial  connections  is  always  a  conspicuous  feature.  With 
these  six  species  in  hand  their  generic  position  becomes  a  second 
problem. 

The  first  name — that  of  Nautilus — is  of  course  used  in  mollusca. 
The  next  available  name — Siderolites  Lamarck — is,  according  to 
various  authors,  the  same  as  Calcarina,  and  if  so  would  have  to  be 
used  by  rules  of  priority  instead  of  Calcarina  d'Orbigny.  It  is  based, 
however,  upon  a  fossil  species  from  the  chalk  of  Maestricht,  and  a 
study  of  its  structure  is  necessary  to  determine  its  true  relations. 
However,  a  study  of  the  material  from  Maestricht  shows  that 
Siderolites  caldtrapoides  is  the  same  generically  as  the  species  fig- 
ured (pi.  45).  Sections  of  the  fossil  material  also  show  that  the 
characters  of  the  two  are  very  similar.  This,  therefore,  is  not  the 
same  as  Calcarina  d'Orbigny,  and  is  not  the  same  as  Gmelin's 
Nautilus  spenglerij  therefore  this  name  is  not  available  as  a  generic 
name  for  the  latter  species.  Tinoporus  Montfort  is  evidently  largely 
based  as  far  as  figure  and  generic  descriptions  show  on  a  species  of 
Calcarina,  although  the  specific  descriptions  in  places  as  noted  by 
Carpenter  and  others  seems  more  like  the  last  of  our  species  men- 
tioned here.  His  remarks  on  the  color  (p.  148),  "blanche,  flambee 
et  teintee  de  jaune  "  would  seem  more  like  the  last,  as  this  is  often 
yellow  or  even  orange  colored.  It  is  evident  therefore  that  the 
genus  Tinoporus  is  in  a  seriously  mixed  condition.  As  has  been 
shown  the  figure  and  generic  description  evidently  refer  to  a  species 
of  Calcarina  but  whether  sufficiently  clear  to  be  used  is  a  question. 


CALCARINA,  TINOPORUS,  AND  BACULOGYPSINA.  367 

Calcarina  d'Orbigny  is  clearly  understood  and  is  the  first  of  the 
names  that  can  be  used  without  question. 

In  1893  Sacco  erected  the  Genus  Baculogysina  on  account  of  the 
uncertainty  of  identifying  Montfort's  Tinoporus.  Sacco  referred 
to  his  genius  Orbitolina  sphaerulata  Parker  and  Jones,  1860,  which 
is  abundant  in  the  white  calcareous  muds  of  Australia.  This  is  the 
reticulately  marked  species  and  this  gives  a  definite  genus  and 
species  for  that  which  may  be  known  as  Baculogypsina  sphaerulata 
(Parker  and  Jones). 

Giimbel  in  1862  describes  Calcarina  tetraedra  from  the  Eocene 
and  this  is  used  as  a  synonym  of  Baculogypsina  sphaerulata  by  vari- 
ous authors.  However,  a  reference  to  his  figure  shows  the  close  rela- 
tion between  this  and  our  species  on  plate  45,  figures  1-5.  A  study 
of  the  fossil  material  of  Siderolites  calcitrapoides  shows  that 
the  fossil  species  and  the  recent  one  from  the  Philippines  are  ap- 
parently generically  the  same.  They  represent  different  species, 
however,  and  probably  our  recent  Philippine  species  is  the  same,  or 
very  close,  to  that  described  by  Giimbel  from  the  Eocene.  If  this 
is  correct,  the  following  key  may  be  used  for  our  six  species.  The 
occurrence  of  Siderolites  as  a  recent  Philippine  genus  is  in  line  with 
that  of  other  groups  of  animals  which  have  now  living  in  this  region 
species  of  genera  elsewhere  extinct. 

A.  Test  rotaliform  throughout ;  with  peripheral  spines,  and  a  supplementary 

skeleton Calcarina    d'Orbigny. 

a1.  Test  fairly  smooth,  spines  smooth  or  channeled. 

ft1.  With  few  spines C.  spengleri  (Gmelin). 

6*.  With  numerous  spines C.  defrancii  d'Orbigny. 

cj.  Test  hispid,  spines  hispid. 

&1.  Flattened,  very  hispid,  spines  numerous C.  hispida  H.  B.  Brady. 

ft2.  Biconvex,    hispid,    centrally    tuberculate,    spines    four    or    five,    dis- 
tinct  C.    baculatus    (Montfort)  ? 

B.  Test  with  rotaliform  young,  hispid,  later  irregular  with   numerous  large 

chambers,  coarsely  punctuate,  usually  three  or  four  blunt  spines,  hispid 

(Siderolites  Lamarck. 

in  the  young,  reticulations  not  distinct •{„.,       ,..  ,       «...., 

(Siderohtes  tetraedra  Giimbel. 

C.  Test  with  very  young  rotaliform,  not  hispid,  later  irregular  with  numerous 

small  finely  punctate  chambers,  4-8  or  more  sharp  spines,  not  hispid, 

(Baculogypsina  Sacco. 

bosses  with  reticulations  very  distinct <  Baculogypsina  sphaerulata 

[  (Parker  and  Jones). 

A  comparison  of  the  distribution  of  Baculogypsina,  sphaerulatus 
(Parker  and  Jones)  and  Siderolites  tetraedra  Giimbel  shows  very 
striking  evidence  of  a  distinct  distribution  for  each.  In  the  Phil- 
ippine region  Siderolites  tetraedra  is  widely  distributed,  and  at  some 
stations  very  abundant,  while  Baculogypsina  sphaerulatus  has  been 
noted  at  but  three  stations  in  the  area,  and  then  as  of  rare  occurrences. 
In  material  from  the  Murray  Island  region  of  the  Great  Barrier  Reef 


368  BULLETIN  100,  UNITED  STATES  NATIONAL  MUSEUM. 

of  Australia  there  is  a  great  development  of  Baculogypsina  sphaeru- 
latus  as  shown  in  the  series  of  samples  collected  by  Dr.  Alfred  G. 
Mayor  on  the  Carnegie  Expedition  to  Murray  Island.  This  species 
makes  up  a  large  proportion  of  the  material  and  Siderolites  is  en- 
tirely lacking.  Baculogypsina  sphaerulatus  is  abundant  at  the  Atoll 
in  Funafuti  according  to  Chapman,  but  no  figures  or  mention  is  made 
of  the  other.  The  more  detailed  distribution  of  these  two  genera  and 
their  relations  to  the  Eocene  would  be  interesting. 

From  their  development,  Calcarina,  with  its  close-coiled  test,  rep- 
resents the  simplest  form,  and  in  its  geological  history  apparently 
goes  back  to  the  Cretaceous.  Siderolites  is  the  next  in  order,  the 
close-coiled  spiral  development  being  continued  for  sometime  in  the 
young  before  the  later  method  of  growth  is  attained.  This  goes 
back  geologically  to  the  Cretaceous.  Baculogypsina  as  applied  here 
has  a  very  short-coiled  stage  in  the  very  young  and  then  assumes  the 
generic  character.  Thus  a  much  more  accelerated  development  is 
shown  and  a  higher  position  in  the  scale.  The  supplementary  skele- 
ton of  bosses  and  radial  connections  is  also  carried  to  a  high  develop- 
ment, making  a  firm  test,  whereas  Siderolites  is  fragile  as  far  as  the 
chambers  are  concerned  (pi.  45,  fig.  4) . 

EXPLANATION    OF    PLATES. 
PLATB  44. 

FIG.  1.  Calcarina  spengleri  (Gmelin)  ventral  view.     X  20. 

2.  Calcarina  defrancii  d'Orbigny  ventral  view  X  20. 

3.  Calcarina  baculatus  ( ?)    (Montfort)  ventral  view.     X  20. 

4.  Calcarina  hispida  H.  B.  Brady  ventral  view.     X  20. 

5.  Siderolites  tetraedra  (Giimbel).     X  15. 

6.  Baculogypsina  sphaerulatus   (Parker  and  Jones).     X  20. 

PLATE  45. 

PIGS.  1-5.  Siderolites  (?)  tetraedra  (Giimbel). 

1.  Young, -ventral  view.     X  45. 

2.  Later  stage,  chambers  invading  dorsal  side.     X  40. 
a,  ventral  side ;  6,  dorsal  side. 

3.  Still  later  stage  where  dorsal  side  is  partly  covered  by  the  invading  chambers. 

X  35. 
o,  ventral  side;  b,  dorsal  side. 

4.  Old  eroded  specimen,  in  which  the  chambers  are  largely  broken  away  leaving 

the  raised  bosses   (x)   as  projections  from  the  center.     X  18. 

5.  Specimens  in  which  the  chambers  now  cover  even  the  spines.     X  15. 

O 


U.  S.   NATIONAL  MUSEUM 


BULLETIN    100     PL.  44 


THE  RELATIONSHIPS  OF  CALCARINA,  TINOPORUS.  AND  BACULOGYPSINA. 

FOR  EXPLANATION  OF  PLATE  SEE  PAGE  368. 


U.  S.   NATIONAL  MUSEUM 


BULLETIN    100     PL.  45 


THE  RELATIONSHIPS  OF  CALCARINA,  TINOPORUS,  AND  BACULOGYPSINA. 


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